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Received: 4 January 2022 Revised: 8 April 2022 Accepted: 13 May 2022
DOI: 10.1111/csp2.12743

CONTRIBUTED PAPER

Human impacts on mammals in and around a protected

area before, during, and after COVID-19 lockdowns

Michael Procko1 | Robin Naidoo2,3 | Valerie LeMay1 | A. Cole Burton1

1
Department of Forest Resources
Management, Forest Sciences Centre,
University of British Columbia,
Vancouver, British Columbia, Canada
2
WWF-US, Washington, District of
Columbia, USA
3 senting a knowledge gap that hinders evidence-based decision-making. We
Institute for Resources, Environment and
Sustainability, University of British
Columbia, Vancouver, British Columbia,
Canada
Correspondence
Michael Procko, Department of Forest
Resources Management, Forest Sciences
Centre, University of British Columbia,
2424 Main Mall, Vancouver, British
Columbia V6T 1Z4, Canada.
Email: xprockox@gmail.com
Funding information
British Columbia Ministry of Environment
and Climate Change Strategy; Canada
Research Chairs program; Natural
Sciences and Engineering Council of
Canada; University of British Columbia
Abstract
The dual mandate for many protected areas (PAs) to simultaneously promote
recreation and conserve biodiversity may be hampered by negative effects of
recreation on wildlife. However, reports of these effects are not consistent, pre-
used camera traps to monitor human activity and terrestrial mammals in
Golden Ears Provincial Park and the adjacent University of British Columbia
Malcolm Knapp Research Forest near Vancouver, Canada, with the objective
of discerning relative effects of various forms of recreation on cougars (Puma
concolor), black bears (Ursus americanus), black-tailed deer (Odocoileus
hemionus), snowshoe hares (Lepus americanus), coyotes (Canis latrans), and
bobcats (Lynx rufus). Additionally, public closures of the study area associated
with the COVD-19 pandemic offered an unprecedented period of human-
exclusion through which to explore these effects. Using Bayesian generalized
mixed-effects models, we detected negative effects of hikers (mean posterior
estimate = 0.58, 95% credible interval [CI] 1.09 to 0.12) on weekly bobcat
habitat use and negative effects of motorized vehicles (estimate = 0.28, 95%
CI 0.61 to 0.05) on weekly black bear habitat use. We also found increased
cougar detection rates in the PA during the COVID-19 closure (esti-
mate = 0.007, 95% CI 0.005 to 0.009), but decreased cougar detection rates
(estimate = 0.006, 95% CI 0.009 to 0.003) and increased black-tailed deer
detection rates (estimate = 0.014, 95% CI 0.002 to 0.026) upon reopening of the
PA. Our results emphasize that effects of human activity on wildlife habitat
use and movement may be species- and/or activity-dependent, and that cam-
era traps can be an invaluable tool for monitoring both wildlife and human
activity, collecting data even when public access is barred. Further, we encour-
age PA managers seeking to promote both biodiversity conservation and recre-
ation to explicitly assess trade-offs between these two goals in their PAs.

KEYWORDS
anthropause, coastal forests, mammal conservation, mesopredator release, predator shield,
recreation ecology

This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided
the original work is properly cited.
© 2022 The Authors. Conservation Science and Practice published by Wiley Periodicals LLC on behalf of Society for Conservation Biology.

Conservation Science and Practice. 2022;4:e12743. wileyonlinelibrary.com/journal/csp2 1 of 15

https://doi.org/10.1111/csp2.12743

2 of 15
1 | INTRODUCTION
Worldwide, the creation of protected areas (PAs) is one
of the most common methods of conserving biodiversity
(Barnes et al., 2017; Sarmento & Berger, 2017), with
many PAs regarded as the only remaining barrier keep-
ing hundreds of species from extinction (Pacifici et al.,
2020). However, PAs are often established under a dual
mandate to conserve biodiversity while also promoting
recreational opportunities, presenting a challenge to PA
managers, as the two may sometimes be at odds
(Sarmento & Berger, 2017; Thomas & Reed, 2019). It is
well known that inadequately regulated consumptive rec-
reation (e.g., hunting, trapping, fishing) may pose a
threat to biodiversity conservation through the direct
removal of wildlife from populations (Schipper et al.,
2008). Nonconsumptive recreational activities (e.g.,
hiking, biking, off-road vehicles) are often thought to
have fewer negative effects on wildlife (Kays et al., 2017),
but evidence suggests they may also hamper PA effective-
ness by imposing widespread disturbance on the wildlife
assemblages inhabiting these spaces (Larson et al., 2016;
Reed & Merenlender, 2008).
Effects of nonconsumptive recreation on wildlife may
include wildlife avoidance of hikers (Erb et al., 2012) and
mountain-bikers (Naidoo & Burton, 2020), increased
physiological stress (Arlettaz et al., 2007), reduced repro-
ductive success (Finney et al., 2005), or increased habitua-
tion leading to shifts in predator–prey dynamics (Geffroy
et al., 2015). To this end, recreation may lead to predators
avoiding areas of higher human influence, while prey spe-
cies and mesopredators may subsequently select these
“predator shields” as a release from top-down pressures
(Berger, 2007; Erb et al., 2012; Prugh et al., 2009;
Sarmento & Berger, 2017). Nevertheless, species exposure
to recreation may vary with park accessibility by people
(Larson et al., 2018) as well as with habitat quality (Reilly
et al., 2017), and methods of assessing recreation effects
on wildlife are sometimes debated (Bateman & Fleming,
2017) or under development (Marion et al., 2020).
In North America, the rapid and continued growth of
the recreation industry (White et al., 2016) reinforces an
urgent need to understand how wildlife respond to larger
numbers of recreationists in PAs. Recreation is well-
known to promote human health (Willis, 2015; Wolf
et al., 2020), cultural resilience (ISRC & CPRA, 2015),
and economic prosperity (Naidoo et al., 2016), and
increases in recreational activity may be driven by popu-
lation growth (White et al., 2016), social media (Winter
et al., 2019), or prolonged summers caused by climate
change (Hewer & Gough, 2018). These increases are com-
monly seen in national parks, but locally managed parks
also see increasing numbers of recreationists, potentially
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PROCKO ET AL.
due to greater accessibility (Larson et al., 2018; White
et al., 2016), underscoring a need for recreation ecologists
to consider how recreation affects wildlife across a range
of PAs.
The British Columbia (BC), Canada provincial park
system is the largest sub-national park system in North
America, with annual visitation estimated at 25.8 million
visitors, and the most heavily used parks each seeing
nearly 1 million day-users (BC Ministry of Environment,
2019a). Over 75% of BC Parks are “dedicated to the pres-
ervation of their natural environments for the inspira-
tion, use and enjoyment of the public” (BC Ministry of
Environment, 2021), with some seeing visitation
increases of over 168% in the last decade (e.g., Joffre
Lakes Provincial Park; BC Ministry of Environment,
2019b). Moreover, BC has the highest levels of biodiver-
sity and species at risk in all of Canada, and is particu-
larly important for larger mammal species (Shackelford
et al., 2018; Westwood et al., 2019), suggesting PA man-
agement and conservation efforts need to be guided by
strategic planning and prioritization (Martin et al., 2018).
However, such evidence-based decision-making may be
difficult without understanding how visitation affects
wildlife in PAs.
Recent lockdowns associated with the COVID-19
pandemic, which some have termed the “anthropause”
(Rutz et al., 2020), provided an unprecedented “experi-
mental” exclusion of human activity which could help
inform how wildlife are impacted by human activity
(Bates et al., 2020). Worldwide, biologists hypothesized
how the natural world might respond to the anthropause
(Corlett et al., 2020; Gaynor et al., 2020; Rutz et al.,
2020), with some of the first emerging studies showing
increased sightings of elusive species (Silva-Rodríguez
et al., 2020) and species reclaiming spaces that had previ-
ously been monopolized by humans (Derryberry et al.,
2020; Manenti et al., 2020; Wilmers et al., 2021). Yet,
these studies have largely focused on urban areas
(e.g., Manenti et al., 2020; Silva-Rodríguez et al., 2020),
exurban residential areas (e.g., Wilmers et al., 2021), or
non-mammalian species (e.g., Derryberry et al., 2020).
Due to the recentness of COVID-19 lockdowns, and the
ongoing nature of the pandemic, there has been little evi-
dence regarding how a lack of recreation during the
anthropause may have impacted mammals in PAs. Fur-
ther, data loss during this time was notable due to many
researchers being unable to safely continue field research
(Pennisi, 2020). This unexpected obstacle highlights the
utility of autonomous monitoring devices like camera
traps (CTs) (Blount et al., 2021), which offer a cost-
effective and non-invasive method of collecting continu-
ous data on both human and wildlife activities (Naidoo &
Burton, 2020).

PROCKO ET AL.
Here, we used CTs to investigate the effects of non-
consumptive recreation on wildlife in a PA and adjacent
research forest at the doorstep of Canada's most densely
populated city—Vancouver, BC (Statistics Canada, 2016).
Our study took place within the context of the COVID-
19-induced anthropause, which mandated a system-wide
shutdown of BC provincial parks for over a month in
spring 2020 and a similar shut-down within the research
forest used in this study. Our objectives were to: (1) quan-
tify the amount and types of anthropogenic activity
occurring throughout this multiple-use landscape; (2) test
the effects of recreational activities on the habitat use of
species inhabiting the area; and (3) investigate how
detection rates of each of these species may have changed
during and after the anthropause. We hypothesized that
larger predatory species such as cougars would be nega-
tively impacted by recreation, implying a form of preda-
tor avoidance of human-derived risk, and that prey
species such as black-tailed deer would therefore be posi-
tively associated with recreation, consistent with the
predator shield hypothesis (Berger, 2007; Sarmento &
Berger, 2017). We also hypothesized that mesopredators
such as coyotes would be positively associated with
human activity, due to the mesopredator release hypothe-
sis (Erb et al., 2012; Prugh et al., 2009). Specifically, we
predicted that while controlling for environmental varia-
tion, areas or times of lower human use would see greater
predator habitat use, and lower prey and mesopredator
habitat use, with areas or times of higher human use see-
ing opposite trends. As a result of these expectations, we
predicted that predator detection rates would increase
during the anthropause, while prey and mesopredator
detection rates would decrease due to anticipated reduc-
tions in human activity during this time.
2 | MATERIALS AND METHODS
2.1 | Study area
This study was conducted in the adjacent landscapes of
Golden Ears Provincial Park (hereafter, “Golden Ears”)
and the University of British Columbia (UBC) Malcolm
Knapp Research Forest (hereafter, “Malcolm Knapp”) in
southwestern BC, Canada (Figure 1). Adjacent to the
Greater Vancouver Region, Golden Ears spans approxi-
mately 625 km2 and supports a range of forest communi-
ties, including lower elevation forests with understories
of vine maple (Acer circinatum) and salmonberry (Rubus
spectaculus) beneath canopies of western hemlock (Tsuga
heterophylla), yellow cedar (Chamaecyparis nootkatensis),
western red cedar (Thuja plicata), and Coastal Douglas-
fir (Pseudotsuga menziesii var. menziewii), and high
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3 of 15
elevation forests with sparse krummholz trees alongside
white mountain-heather, mosses, and lichens
(BC Ministry of Environment, 2013). The park permits a
variety of recreational activities such as camping, hiking,
mountain biking, horseback riding, fishing and boating,
but excludes hunting and off-road vehicles. Golden Ears
also contains the largest campground in the province,
receives almost one million recreational visitors annually,
and recently reported the most annual campers and the
second most annual day-users of any BC provincial park
(BC Ministry of Environment, 2019a). These recreational
pressures are the main form of anthropogenic disturbance
in the park, as resource extraction has been prohibited
since 1929, rendering the park a prime example of a
protected coastal forest with high levels of human
visitation.
Malcolm Knapp is a 52 km2 research forest that abuts
the southwestern boundary of Golden Ears and contains a
similar forest ecosystem, but has the principal management
objectives of providing research and educational opportuni-
ties for UBC students and others. Designated as a research
forest reserve in 1943, the privately owned forest includes
an extensive network of over 100 km of roads, which facili-
tate the main goals of education and research activities, as
well as forestry operations (e.g., harvesting, thinning, fertil-
ization, and replanting). Malcolm Knapp also receives
thousands of recreational visitors annually who access
approximately 30 km of trails. However, road access is
curtailed via a strict gated access route, and camping, hunt-
ing, fishing, and off-road recreation are not permitted in
the forest. There are also reservable facilities for recreation
and other user groups outside of UBC student groups, but
activities associated with these facilities are contained
within a small (<2 km2) portion of the forest. Roads in
Malcolm Knapp can be used for hiking, but motorized
usage is restricted only to those with access to the main
gate (i.e., Malcolm Knapp staff, UBC researchers, and
groups registered for activities on site). Therefore, the main
drivers of human disturbance in Malcolm Knapp are
related to educational and research activities, including for-
estry operations, but some areas see mild-to-moderate
levels of nonmotorized day-use recreation.
In the Spring of 2020, Golden Ears and Malcolm Knapp
were temporarily closed due to the COVID-19 pandemic.
Golden Ears was closed to the public from 8 April to May
13, 2020, after which it opened for limited day-use (under a
permit system) on 14 May and camping on June 1, 2020.
Malcolm Knapp was closed from 23 March to June 5, 2020.
During these times, public access was heavily restricted,
with substantial fines being imposed on individuals who
entered these areas without authorization. However, lim-
ited education, research, and forest operations in Malcolm
Knapp were permitted during the closure.
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4 of 15 PROCKO ET AL.

F I G U R E 1 Study area of Golden Ears Provincial Park (red outline) and Malcolm Knapp Research Forest (orange outline) in
southwestern British Columbia, Canada. Black circles represent on-trail camera trap locations, and white circles represent off-trail camera
trap locations. Black lines represent roads, brown lines represent recreational trails, green (hashed) polygons represent agricultural land
reserves, red polygons represent harvested forest from the past 5–20 years, and blue polygons/lines represent hydrological features (lakes/
streams)

2.2 | Camera trap sampling design
Beginning in March 2019, we deployed a stratified ran-
dom CT array both on and off human-use trails and
roads to monitor medium- and large-bodied mammals,
hikers, mountain bikers, horseback riders, and vehicles.
Strata in Golden Ears included locations along high
human use trails, low human use trails, and off-trail
(cameras placed >250 m and <1 km from trails). High/
low use designations of trails were based on local knowl-
edge and expert opinion of park managers (S. Stickney,
pers. comm). In Malcolm Knapp, two strata were used:

PROCKO ET AL.
on-trail (or road) and off-trail, with the latter similarly
having cameras placed >250 m and <1 km from both
trails and roads. We used a buffer of 250 m to 1 km due
to difficulty accessing off-trail locations in the steep park
terrain. Cameras were separated by a minimum of 300 m.
We navigated as close as feasible to each random
point and attached a Reconyx Hyperfire Pro 2 (Reconyx,
Holmen, WI, USA) CT to a nearby tree. CTs were set for
a target detection zone, focused on either a recreation
trail or road (“on-trail”), or a game trail (“off-trail”)
approximately 3–5 m in front of the lens. CTs were set at
approximately 1 m in height to maximize detection prob-
ability of medium- and large-bodied wildlife species,
while also restricting the amount of identifiable human
features captured in photos to protect the privacy of indi-
viduals using the landscape. We took care to ensure cam-
era heights and distances to target zones were similar
across the study, as small changes in these can drastically
impact detectability (Miller et al., 2017), thereby biasing
results.
The final array consisted of n = 58 camera “stations,”
including 37 in Golden Ears (20 on-trail, 17 off) and 21 in
Malcolm Knapp (13 on-trail or road, and 8 off). Most sta-
tions were active for at least 1 year, with some periods of
camera inactivity due to malfunctions, vandalism, and
theft (Appendix S1), but periodic SD card collections
were performed to minimize these until final collections
in August–September 2020.
Photos containing humans were blurred using a facial
redaction software (Lixar, 2018) to ensure privacy protec-
tion. These images were then uploaded to a custom cam-
era trapping database for identification (WildCo, 2021),
which utilized an automated detection software to sort
the photos into human, vehicle, and animal categories,
before subsequent review by the project team (Fennell
et al., 2022). We identified species for all images con-
taining mammalian wildlife and domestic dogs, and we
classified images of humans into the following categories:
hikers, mountain bikers, horseback riders, and motorized
vehicles. Data were then imported to R (version 3.6.2) for
data management and statistical analyses (R Core
team, 2019).
2.3 | Analytical framework
Prior to modeling, we defined independent detections of
wildlife as images of the same species taken at least 30
min apart at each station (Burton et al., 2015). Indepen-
dent detections of human activity were determined using
an independence threshold of 1 min, since numerous rec-
reationists were likely to use high-traffic trails within the
span of 30 min. We therefore assumed that multiple
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5 of 15
wildlife photos taken within 30 min of each other con-
tained the same individuals or groups, while multiple
human photos taken more than 1 min after another
would contain different individuals or groups, given most
recreationists travel unidirectionally along trails. We
selected focal wildlife species for modeling using three
criteria: (1) those with at least 30 independent detections;
(2) those with at least medium body size (>1 kg); and
(3) those with primarily terrestrial rather than arboreal
movement. Our response variable was a binary detec-
tion/non-detection for each focal species during each
week at each station. We chose this “station-week” tem-
poral scale to assess temporal variation in species habitat
use in response to human activity, while avoiding an
excess of zeros in detection histories at finer temporal
scales (e.g., “station-days”; Naidoo & Burton, 2020).
To investigate how recreational activities impacted
wildlife habitat use, we modeled the “probability of use”
(Kays et al., 2021) for each of these focal species during a
station-week with a Bayesian generalized linear mixed-
effects model (GLMM), assuming a binomial response
distribution and including station as a random intercept
to account for non-independence among weeks sampled
at the same station. Using this GLMM specification, the
probability of use of a station during each week for each
species was modeled as a function of several hypothe-
sized explanatory variables (Table 1). Since our primary
focus was on species responses to recreational activities,
we investigated possible effects of hikers, mounted recre-
ationists (i.e., horseback riders and mountain bikers),
and motorized vehicles during a given station-week by
including the average weekly detection rates
(i.e., number of detections at a station-week divided by
the number of days the camera was active during that
week, typically 7) as possible explanatory variables.
Eleven additional variables were included to control
for alternative hypothesized drivers of wildlife habitat use
variability. These included forest measures, namely,
crown closure (%), forest height (m), the percent of forest
recently (within 5–20 years) harvested within a 500 m
buffer of the station (%), and topographical measures,
specifically elevation (m) and slope (angle in  ). We con-
trolled for seasonal variation in vegetation productivity
using the normalized difference vegetation index (NDVI)
calculated using MODIS product VNP13A1 (8-day inter-
val data at 500 m resolution). We also included distance
to nearest water source (m) as a general habitat measure,
and distance to the urban-wildland boundary (m) as an
index of the potential influence of residential and agricul-
tural areas surrounding the PA. Finally, we included vari-
ables describing variation in deployment across stations
that could affect detectability (Hofmeester et al., 2018),
namely the height of the camera above the intended

6 of 15
target path (m above the recreation trail, road, or game
trail), the distance to the intended target path (m), and
whether the camera was placed along a road or trail
(binary on/off). We therefore assumed that the potential
influence of variation in detectability was minimized by
our standardized protocols and inclusion of these sam-
pling variables. We did not use an occupancy modeling
approach to estimate detectability, as we were interested
in variation in detections as a signal of local habitat use
rather than as detection error, and we considered it
unlikely that the assumptions of occupancy modeling
would be met in our sampling context (e.g., site closure;
Neilson et al., 2018; Kays et al., 2021). We assumed any
temporal autocorrelation was accounted for through the
inclusion of the random effect of camera station, and
tested for spatial autocorrelation among stations using
Mantel tests on model residuals.
To test whether the anthropause impacted focal spe-
cies, we built three additional Bayesian models for each
species utilizing data from: (1) the whole study area;
(2) only Golden Ears; and (3) only Malcolm Knapp. We
included all dates during the closure, but only pre-closure
(2019) data matching the sample dates for the post-
closure (2020) period (early-May to mid-September) were
included, to reduce the possibility of any additional
weight toward pre-closure trends given the longer pre-
closure sampling period. We also calculated detection
rates (i.e., the number of detections at a station divided
by number of days the station was active rather than raw
detections) to account for the different sampling effort
within the shorter period of the anthropause relative to
the longer pre- and post-closure periods. Each model
predicted the detection rate of a species before, during,
and after the closure time periods using a factor variable
to alter the intercept by time period. We used the closure
period as the baseline intercept, and tested whether wild-
life detection rates before or after the closure differed
from the closure period. We did not include other explan-
atory variables, such as habitat type, as these were consis-
tent for each station throughout the three comparison
periods given stations were not moved and time periods
represented similar seasons.
For the models used to examine recreational activity
effects on probability of habitat use, we tested for exces-
sive multicollinearity that would cause parameter estima-
tion instability using pairwise correlations (all r < 0.7;
Zuur et al., 2010) (Appendix S2). We used Kendall's tau
and Spearman's rho for any correlations with the bino-
mial variable, Trail, and Pearson's correlation for all
other pairs. We normalized all continuous explanatory
variables by subtracting the mean and dividing by the
standard deviation (SD) to facilitate comparisons of rela-
tive effects of variables on the probability of species
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PROCKO ET AL.
habitat use. These models were constructed using flat
priors, and run with four chains of 100,000 iterations
(burn-in period of 50,000) to ensure model convergence.
Models to investigate changes in detection rates between
time periods were constructed with flat priors, and run
with four shorter chains of 10,000 iterations (burn-in
period of 2000), which were sufficient for model conver-
gence. For all models, convergence of posterior distribu-
tions was confirmed visually with trace plots and with
the Gelman-Rubin statistic (R-hat) (Hobbs & Hooten,
2015). Variables were considered to have strong evidence
for effects on wildlife detections if 0 was not within their
95% credible parameter estimate intervals. Models were
implemented using the R package brms (Burkner, 2017).
3 | RESULTS
3.1 | Summary of survey
From the 58 CTs deployed across Golden Ears and Mal-
colm Knapp, we collected and classified 1,059,703 images
from 23,928 camera trap-days of sampling effort, which
provided detection data for 3479 station-weeks used in
modeling. We detected 19 mammalian wildlife species
(Appendix S3), six of which met our criteria for modeling,
namely, black-tailed deer (Odocoileus hemionus, n = 709
independent detections using 30 min. threshold), coyote
(Canis latrans, n = 416), black bear (Ursus americanus,
n = 290), snowshoe hare (Lepus americanus, n = 248),
bobcat (Lynx rufus, n = 203), and cougar (Puma concolor,
n = 46). Black bears were the most widely distributed of
these wildlife species, being detected at 44 of our 58 sta-
tions (76%), whereas snowshoe hares showed the most
restricted distribution of our six focal species, being
detected at only 26 (45%) of all stations (Appendix S3).
Notable detections of species that were too infrequently
detected to model included the possibly threatened but
data deficient western spotted skunk (Spilogale gracilis,
n = 7) and one individual from a recently (c. 2007–2008)
reintroduced elk population (Roosevelt subspecies;
Cervus canadensis roosevelti, n = 1).
Our camera-trap sampling also collected over 111,486
independent detections of humans (1 min. threshold).
Detected activities included 108,865 hiker detections,
1584 mounted recreationists, and 1037 vehicles, with a
majority of hikers (n = 104,325) and all but one mounted
recreationist (n = 1583) being detected in Golden Ears,
and nearly all vehicles (n = 946) being detected on roads
in Malcolm Knapp. Although we also identified 31,562
detections of domestic dogs, these were highly correlated
with hikers (r = 0.92) and therefore not used as a sepa-
rate predictor variable in models. Park closures resulting
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PROCKO ET AL. 7 of 15

T A B L E 1 Predictor variables used in the Bayesian generalized linear mixed-effects models, which contrasted the probability of weekly
habitat use for focal species against measures of various forms of human activity while controlling for alternative sources of variation. The
mean, minimum (min), and maximum (max) of values (on the raw scale) are provided for the 58 camera stations

Variable Description Acquisition Hypothesis Mean Min Max

a
Hikers Weekly detection rate per station- CT Negative impact on top predators, 4.68 0.00 191.50
week (# hikers detected per week/ positive impact on prey and
# days the camera was active that mesopredators (predator shield &
week) mesopredator release)
Mounted Weekly detection rate per station- CT Negative impact on top predators, 0.07 0.00 6.00
recreation week (# mountain bikers detected positive impact on prey and
per week/# days the camera was mesopredators (predator shield &
active that week) mesopredator release)
Motorized Weekly detection rate per station- CT Negative impact on top predators, 0.04 0.00 9.29
vehicles week (# vehicles detected per positive impact on prey and
week/# days the camera was active mesopredators (predator shield &
that week) mesopredator release)
Crown closureb % crown closure at site GISc Control for habitat type 64.47 10.00 85.00
b
Stand height Projected height of forest at site (m) GIS Control for habitat type 33.03 12.10 53.30
NDVId Normalized difference vegetation GIS Control for seasonality 0.75 0.35 0.99
index in a week at site, measured
in 8-day intervals (500 m
resolution)
Pct. Harvested % of recently (between 2000 and GIS Control for habitat type and resource 0.06 0.00 0.37
2015) harvested forest within a 500 availability
m buffer around each station
Distance to Distance to the nearest stream, river, GIS Control for resource availability 152.94 3.90 744.17
watere or lake from site (m)
Distance to south Distance to the nearest urban- GIS Control for influence of residential 4466.29 39.97 17,008.14
boundaryf wildlands boundary (m) areas adjacent to PA
Elevationg Elevation at site (m) (25 m GIS Control for topography 338.29 11.00 1150.00
resolution)
Slopeg Slope at site (degrees) (25 m GIS Control for topography 13.38 1.28 39.92
resolution)
Trail Binary indication of whether site was Fieldh Control for camera set 0.58 0.00 1.00
on-trail/road or off
Camera height Height (m) the camera was position Field Control for camera set 0.76 0.41 1.24
at each site
Distance to target Distance from the camera lens to the Field Control for camera set 3.46 1.32 7.54
anticipated path of the target
(either the center of the trail/road
or the nearest game trail) (m)
a
CT acquisition method—data were collected by camera trap.
b
Data Source: Data Management and Access—Province of British Columbia.
c
GIS acquisition method—data were collected using geoprocessing tools in ArcGIS Pro.
d
Data Source: MODIS Satellite Product VNP13A1.
e
Data Source: Freshwater Atlas—Province of British Columbia.
f
Data Source: Shapefiles of boundaries provided by BC Parks and Malcolm Knapp Research Forest Management.
g
Data Source: Digital Elevation Model—Province of British Columbia.
h
Field acquisition method—data were collected in the field.

from the COVID-19 pandemic were very effective, lead- maintenance, forest harvest) were detected during the
ing to reduced recreationist detections during these times closure at comparable numbers to pre- or post-closure
(Figure 2). However, working operations (e.g., park (Figure 2).
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8 of 15 PROCKO ET AL.

F I G U R E 2 Weekly detections per active camera of (a) hikers, (b) mountain bikers, (c) horseback riders, and (d) motorized vehicles.
Horizontal axes represent time throughout the year (in weeks), and vertical axes represent the number of detections per active camera
(standardized rate to account for differences in sampling effort among weeks). Spaces between the red dashed lines indicate the COVID-
19-related closure periods of Golden ears Provincial Park, and spaces between the blue dashed lines indicate the COVID-19-related closure
periods of Malcolm Knapp Research Forest

3.2 | Drivers of weekly wildlife
habitat use
Of the habitat use models run for the six focal species,
two showed strong evidence for associations between
wildlife and human activity (all models successfully con-
verged with R-hat <1.1 for all model terms; Appendix
S4). Specifically, bobcat habitat use was negatively associ-
ated with hikers (mean posterior estimate = 0.58, 95%
credible interval [CI] 1.09 to 0.12; Figure 3) and black
bears were negatively associated with motorized vehicles
(estimate = 0.28, 95% CI 0.61 to 0.05; Figure 3). No
other species showed strong evidence of associations with
direct measures of human activity. However, several
other predictor variables included to control for variation
showed strong evidence of impacting species' probability
of habitat use (Figure 3). We identified strong evidence
that the distance from each station to the urban-wildland
boundary influenced the probability of habitat use for
four of six species (all except black bears and cougars),
with black-tailed deer, snowshoe hares, coyotes, and bob-
cats all being more likely to be detected at stations closer
to the boundary (estimates & 95% CIs: black-tailed deer
= 0.87, 1.65 to 0.12, snowshoe hares = 1.82, 3.44
to 0.51, coyotes = 1.57, 2.73 to 0.60, bobcats =
1.19, 2.09 to 0.37). Whether the camera was situated
along a trail or road was positively associated with proba-
bility of habitat use for all predatory species (estimate &
95% CI: cougars = 2.04, 1.11 to 3.10, black bears = 1.38,
0.59 to 2.21, coyotes = 2.69, 1.46 to 4.11, bobcats = 3.36,
2.11 to 4.74). Further, environmental variation indicated
by variables such as elevation, slope, and NDVI also
showed strong evidence of impacting species' habitat use
(Figure 3).
3.3 | Wildlife detection rates
in the Anthropause
From models constructed with data from the entire study
area, cougars were the only species to show greater detec-
tion rates during the anthropause (estimate = 0.005, 95%
CI 0.004 to 0.007, Appendix S5) than before or after the
anthropause, with the intercept of the detection model
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PROCKO ET AL. 9 of 15

F I G U R E 3 Predictor variables
(y-axis) which were strongly associated
with species probability of habitat use in
Bayesian generalized linear models
contrasting species probability of habitat
use against a suite of explanatory
variables. Focal species (from top to
bottom) included cougar, black bear,
black-tailed deer, snowshoe hare, coyote,
and bobcat. The horizontal (x) axis
illustrates posterior distributions, with
points in each line representing mean
posterior estimates and blue lines
representing 95% credible intervals. The
hashed green line is the x-intercept of
0, which was used to determine strength
of predictor variables (predictor
variables had strong evidence if 95%
credible intervals of posterior estimates
did not overlap zero)

F I G U R E 4 Detection rates
(detections/camera days; y-axes) per
station, with data restricted to Golden
Ears, before, during, and after the
COVID-19-related closures (x-axis), for
cougars (blue dots and lines; left y-axis)
and black-tailed deer (red dots and lines;
right y-axis). Each dot represents the
detection rate for a single camera during
a given period, with connecting lines
illustrating changes between periods.
Also shown are the means with
whiskers showing the standard errors
around these means

dropping by 0.005 for before (estimate = 0.005, 95%
CI 0.007 to 0.003) or by 0.004 after (estimate =
0.004, 95% CI 0.006 to 0.002) relative to during the
anthropause. When data were restricted to Golden Ears,
cougars were similarly detected at a higher rate during
the closure (estimate = 0.007, 95% CI 0.005 to 0.009) than
before or after the closure (both estimates = 0.006, both
95% CIs 0.009 to 0.003), while black-tailed deer
showed higher detection rates after the closure (estimate
= 0.014, 95% CI 0.002 to 0.026) than during (estimate =
0.002, 95% CI 0.007 to 0.010), but no difference was
found between during and before (Figure 4, Appendix

10 of 15
S5). In Malcolm Knapp, there was no strong evidence of
changes in species detection rates across time periods
(Appendix S5).
4 | DISCUSSION
Our findings provide evidence of shifts in wildlife habitat
use in response to variation in recreational pressure at
the weekly scale, as well as dramatic changes in recrea-
tional activity as a result of the closures imposed by the
COVID-19 pandemic, coinciding with some shifts in spe-
cies' detection rates. We predicted that larger predators
would avoid areas and times of higher recreation, all-
owing prey and mesopredator species to evade predation
pressure by positively associating with recreation. These
predictions were not supported by the data, with mes-
opredator bobcats being negatively impacted by hikers
and large predator black bears being negatively impacted
by motorized vehicles. Negative relationships between
bobcats and hikers have been observed elsewhere
(George & Crooks, 2006; Reed & Merenlender, 2008).
Therefore, while this finding does not support the mes-
opredator release hypothesis, it provides additional sup-
port for concerns that high levels of recreation may
impact bobcat habitat use. Additionally, the strong nega-
tive relationship between black bear detections and
motorized vehicles is interesting in the context of this
study area since most vehicle detections were tied to
Malcolm Knapp research and education activities, as well
as timber harvesting to facilitate these activities. This
implies that although black bears are well-known to
select for regenerating stands following forest harvest
(Brodeur et al., 2008), they likely avoid areas with very
high human activity, particularly where harvest and
other forest management interventions are taking place.
Prior research has confirmed that bears tend to avoid
spaces or times of higher motorized vehicle activity
(Ladle et al., 2018; Naidoo & Burton, 2020), but these
results typically pertain to grizzly bears (Ursus arctos).
Others have shown black bears avoid roads (Carter et al.,
2010; Zeller et al., 2019), but these do not measure motor-
ized disturbance directly, and instead use the presence of
roads as a proxy for motorized disturbance. Research
regarding changes in black bear habitat use in response
to direct measures of vehicle activity is sparse given the
difficulty in collecting data on both. Thus, we present a
novel account of black bear habitat use being negatively
impacted by explicit measurements of vehicle use.
Alternatively, the negative association between the
distance to the urban-wildland boundary and habitat use
of all prey and mesopredator species may provide a
different form of support for the predator shield and
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PROCKO ET AL.
mesopredator release hypotheses, given human activity
and footprint can differentially impact wildlife (Nickel
et al., 2020). Accordingly, prey and mesopredator species
may use residential areas near the PA as a shield from
larger predatory species. Cougars are known to avoid
housing densities above certain limits (Smith et al.,
2019), and although cougars commonly traverse urban-
wildland gradients (Alldredge et al., 2019), cougar mor-
tality is often greater in more developed areas (Moss
et al., 2015). Likewise, black bears commonly associate
with exurban spaces due to anthropogenic resource avail-
ability, but face a similar risk of mortality (Braunstein
et al., 2020; Laufenberg et al., 2018), which could poten-
tially provide a shield for mesopredator or prey species.
Therefore, we suggest future research could target a gra-
dient from exurban residential neighborhoods to PAs to
investigate whether larger predators are less likely to use
spaces adjacent to PAs, and whether prey or mes-
opredators more commonly inhabit these areas to avoid
predators.
We also found strong evidence that cameras situated
along trails or roads were more likely to detect predator
and mesopredator species. Prior work has identified pref-
erences for linear features in several mammalian species
(Fisher & Burton, 2018), even in regions with strong rec-
reational pressures (Reilly et al., 2017). There is thus
potential for animal preference for trails to obscure
responses to human activities. However, our inclusion of
off-trail (i.e., game trail) samples and the corresponding
binary model covariate controlled for variation in detec-
tions due to on- or off-trail placement of CTs. Neverthe-
less, we recommend future research with more extensive
off-trail sampling (e.g., proportional to available off-trail
habitat) to further evaluate potential confounds between
animal trail use and responses to recreation. Addition-
ally, alternative analytical approaches could be used to
assess potential finer-scaled trade-offs for species that use
trails but avoid human activities (e.g., attraction-
avoidance ratios; Naidoo & Burton, 2020).
Our CT sampling provided a unique opportunity to
investigate how wildlife responded to the anthropause
when the study area was closed to the public. Both the
strong increase in cougar detection rates during the
anthropause and the strong increase in black-tailed deer
detection rates following the reopening of Golden Ears
provide support for the predator shield hypothesis in the
context of recreation. Cougars are known to exhibit fear-
responses in response to human voices (Smith et al.,
2017; Suraci et al., 2019), and have elsewhere responded
positively to the anthropause, expanding their ranges into
previously unoccupied areas during this time (Wilmers
et al., 2021). This immediate response from a top predator
to reductions in human activity has implications for

PROCKO ET AL.
carnivore–human coexistence in shared landscapes, unde-
rscoring the ability of carnivores to quickly adapt to human
activity (Carter & Linnell, 2016). Additionally, increases in
black-tailed deer detections during the post-closure period
may have been related to the implementation of a day-use
pass system which limited recreation upon the reopening
of Golden Ears. This implies the possibility of a threshold
in recreation effects, whereby intermediate levels may pro-
vide a predator shield by displacing cougars but not deer,
while higher levels may displace both predators and prey.
Identifying such thresholds of disturbance is integral to
providing science-based recommendations to PA managers
(Dertien et al., 2021). Thus, our findings provide support
for the use of continuous measures of recreation intensity
to inform trail limits. We recommend that future research
consider analytical methods which could be used to assess
thresholds of recreational disturbance (e.g., nonlinear
models: Pinheiro & Bates, 2000; piecewise regression: Malo
et al., 2011, Toms & Lesperance, 2003; additive models:
Zuur et al., 2009), and thus identify recreation “carrying
capacities” to inform park management.
The differences in deer responses to recreation that
we observed at shorter (weekly) vs. longer (park closure)
periods emphasizes the importance of considering tempo-
ral scale in wildlife research. Human–wildlife coexistence
can occur across different scales, with some species facili-
tating coexistence by segregating from people at coarser
temporal scales (e.g., weekly, Naidoo & Burton, 2020),
while other species may avoid humans at finer scales
(e.g., daily, Patten & Burger, 2018, or even hourly, Carter
et al., 2012; Nickel et al., 2020). The weekly temporal
scale of our first set of models did not consider potential
finer-scale segregation, which may have provided more
information regarding wildlife displacement by human
activity. However, given the lack of previous monitoring
in this landscape, we sought to investigate broader pat-
terns of species composition and distribution, while
understanding how these metrics were associated with
coarse-scale anthropogenic (e.g., weekly hiker traffic) or
environmental factors (e.g., NDVI). Likewise, our second
set of models also made coarse-scale comparisons, sum-
marizing how wildlife detection rates changed through-
out time periods of multiple weeks. However, given the
limited closure period and our goal of understanding
how wildlife activity differed between the closure and
periods when the study area was open, we contend that
investigation at this temporal scale was most appropriate.
Future work may harness analytical methods better
suited to investigate whether wildlife segregate from rec-
reationists at daily temporal scales, or whether even
finer-scale behavioral shifts may facilitate human-wildlife
coexistence in this system (e.g., altered diel activity
patterns, Carter et al., 2012).
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11 of 15
5 | CONCLUSIONS
The contrast between a heavily visited protected area and
a less recreated research forest with limited public road
access provided a unique context in which to assess the
effects of anthropogenic disturbance on mammalian hab-
itat use. This, in addition to the restriction of public
access for a number of weeks during the anthropause
offered a rare unplanned experiment through which to
explore effects of human activity on wildlife. We aimed
to test both the predator shield and mesopredator release
hypotheses in the context of recreational activity, finding
little support for either hypothesis at the weekly scale,
but stronger support for the predator shield hypothesis
when considering how detection rates of focal species
varied during and after the anthropause. Specifically, at
the weekly scale, our findings indicated that recreation in
PAs may indeed hamper some wildlife habitat use, with
species such as bobcats being displaced by larger quanti-
ties of hikers. Moreover, forest operations and other
activities in landscapes adjacent to PAs may also nega-
tively impact black bears, resulting in a mosaic of distur-
bances for the species. We also found increases in cougar
detection rates during the anthropause, with the subse-
quent reopening of the PA coinciding with decreased
cougar detection rates and increased black-tailed deer
detection rates. Thus, our study emphasizes how PAs
faced with a challenge to simultaneously conserve wild-
life while promoting recreation may benefit from consid-
ering species- and context-specific approaches to wildlife
management, with a particular emphasis on the most
sensitive species. We also underscore the utility of CTs in
continuously monitoring recreation and wildlife in PAs
both generally, and through periods when public access
to these spaces is restricted. In light of the dual mandate
for many PAs to provide opportunities for recreation
while simultaneously conserving wildlife, we encourage
PA managers worldwide to adopt tools (e.g., CTs, trail
counters) and strategies (e.g., collaboration with external
groups) which could better inform on trends in recrea-
tion and wildlife habitat use in order to understand
whether the two are at odds in their own systems. While
there are undoubtedly financial and logistical constraints
to detailed investigations of these trends in many PAs, we
suggest that targeted monitoring, research partnerships,
and cost-effective protocols can help overcome current
data deficiencies.
AUTHOR CONTRIBUTIONS
A. Cole Burton conceived the study. Michael Procko and
A. Cole Burton conducted the field work. Michael Procko
conducted the data analysis. Michael Procko, Robin
Naidoo, Valerie LeMay, and A. Cole Burton wrote the

12 of 15
paper. A. Cole Burton acquired funding to support the
study.
A C K N O WL E D G M E N T S
This research was supported by the British Columbia
Ministry of Environment and Climate Change Strategy,
the Natural Sciences and Engineering Council of Canada,
and the Canada Research Chairs program. We thank
Mitch Fennell, Taylor Justason, Katie Tjaden-McClement,
Isla Francis, and Gerlissa Chan for help deploying and
checking camera sites; Katie Tjaden-McClement, Isla
Francis, and Gerlissa Chan for help classifying photos;
and Catherine Sun, Chris Beirne, and Alys Granados for
comments on preliminary drafts and analytical consider-
ations. We also thank Melanie Percy, James Quayle,
Sam Stickney, Joanna Hirner, Daris Lapointe, Simon
Debisschop, Gareth Wheatley, and Ionut Aron for assis-
tance and support.
CONFLICT OF INTEREST
The authors declare no conflicts of interest.
DATA AVAILABILITY STATEMENT
Data and materials are available in a publicly accessible
repository: https://doi.org/10.5061/dryad.hx3ffbggk.
Additional information is available online in the
Supporting Information section at the end of the online
article. The authors are solely responsible for the content
and functionality of these materials. Queries (other than
absence of the material) should be directed to the
corresponding author.
E TH IC S ST A T EME N T
This research was conducted under protocol A18-0234
from the University of British Columbia's Animal Care
Committee and under protocol H21-01424 from the Uni-
versity of British Columbia's Behavioral Research Ethics
Board.
ORCID
Michael Procko https://orcid.org/0000-0002-0302-9058
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How to cite this article: Procko, M., Naidoo, R.,
LeMay, V., & Burton, A. C. (2022). Human impacts
on mammals in and around a protected area
before, during, and after COVID-19 lockdowns.
Conservation Science and Practice, 4(7), e12743.
https://doi.org/10.1111/csp2.12743